Caribbean datums and the integration of geographical …webb.deu.edu.tr/sacitozer/kitaplar/201310191306511.pdf · fragmented rudist shells and an upper thinner unit composed of silty

Tunis, G., Özer, S., Radoičić, R., Săsăran, L., Tarlao, A. and Tentor, M. 2013. The state of the knowledge of the

genus Pseudopolyconites. Caribbean Journal of Earth Science, 45, 107-118. © Geological Society of Jamaica.

Available online 14th September 2013.

107

The state of the knowledge of the genus Pseudopolyconites

GIORGIO TUNIS1, SACIT ÖZER

2, RAJKA RADOIČIĆ

3, LIANA SĂSĂRAN

4, ALCEO TARLAO

5

AND MAURIZIO TENTOR5

1Dipartimento di Geoscienze, Università di Trieste, via Weiss 2, 34127 Trieste, Italy. Email: [email protected]

2Dokuz Eylül Üniversitesi Mühendislik Fakültesi Jeoloji Mühendisliği, Bölümü, Tınaztepe Yerleşkesi 35160

Buca Kampusu İzmir, Turkey. 3Kralija Petra, 38-VI, Beograd, Serbija.

4Department of Geology, Babeş-Bolyai University, Str. Kogălniceanu 1, 40084, Cluj-Napoca, Romania.

5Museo della Rocca di Monfalcone, via Valentinis 134, 34074 Monfalcone, Italy.

ABSTRACT. The genus Pseudopolyconites is distributed in Upper Cretaceous successions of the Central Tethys.

Specimens have been recovered from carbonate, siliciclastic, volcanoclastic or mixed carbonate/siliciclastic

successions dated from the late Santonian or early Campanian to the latest Maastrichtian, but often only

fragmented shells have been collected and described. Many species have been established but the constructional

morphology of individuals and, in particular, the characters of the outer shell layer were seldom adequately

examined by rudistologists. Currently two species can be recognized with certainty: Pseudopolyconites hirsutus

(late Santonian-early Campanian) and Pseudopolyconites serbicus (late Campanian). Currently, the phylogenetic

relationships within the genus during the Santonian to latest Maastrichtian remain obscure. It is speculated that

the examples of Pseudopolyconites found at the type-locality of Bačevica developed particular shell growth

strategies in response to a peculiar environment where they thrived.

Key words: Pseudopolyconites; morphology; palaeo-biogeographic distribution; Upper Cretaceous.

1. INTRODUCTION

The rudist bivalve genus Pseudopolyconites was

established in 1934 by the Serbian palaeontologist

Branislav Milovanović (1934a). The first

specimens of this rudist were found close to the

village of Bačevica (eastern Serbia) which is

considered the type-locality of the genus. Due to

the complete disappearance of all the holotypes and

paratypes of the numerous species of

Pseudopolyconites established by the Serbian

rudistologists from Bačevica, we have surveyed

and examined many times the lithological and

faunal succession cropping out in the area in

question. The scope of the latest investigation has

been aimed at collecting examples of

Pseudopolyconites and at examining the faunal

assemblages close to the Pseudopolyconites

bearing strata. Successively we have expanded the

research upon the Pseudopolyconites specimens

found in Rumania and Turkey and we have

examined the literature concerning all known

Pseudopolyconites bearing strata both within

carbonate and clastic successions. In some cases we

have surveyed carbonate successions cropping out

in the Adriatic region.

The aim of this paper is to discuss different

aspects concerning the genus Pseudopolyconites:

that is, morphological characters, faunal

assemblages, age, palaeobiogeographical

distribution, phylogeny and possible developments

of the research.

2. LITHOLOGICAL AND FAUNAL CHARACTERISTICS

OF THE BAČEVICA SUCCESSION

Due to the shortage of outcrops in the environs of

Bačevica (Figure 1) and the low quality of the

exposures, it is only possible to make a brief

assessment of the lithological succession and the

field relationships. The lower part of the Bačevica

succession is characterized by rhythms consisting of

a lower unit of thick limestone breccias containing

fragmented rudist shells and an upper thinner unit

composed of silty limestones with abundant, often

intact rudists. The rhythms record multiple events of

transport and reworking of sediments and of

recolonization by rudists (Tarlao et al., 2010). The

upper part of the Bačevica succession is represented

mainly by weathered sandstones and siltstones with

subordinate conglomerates and silty limestones.

Because of the rare and very limited outcrops and

the shallow dip of the strata, the total thickness of

the section is very difficult to estimate.

In the absence of strontium isotope analyses

(SIS), the inferred age of the upper part of the

Bačevica section is Campanian on the basis of close

similarities with the Gavlo section (see below).

Tunis et al. The state of the knowledge of the genus Pseudopolyconites

107

Figure 1. Geographical map of the vast considered area. The numbers indicate the different localities mentioned

in the text: 1, Remeti valley; 2, Dealul Magura; 3, Garlo; 4, Vrla Strana, Vina; 5, Hron valley; 6, Hvar island; 7,

Mt. Nanos; 8. Anhovo; 9, Maniago; 10, Cava Soframa; 11, Vitagliano; 12, Ciolo cove; 13, Zakynthos island; 14,

Lefkas island; 15, Kocaeli peninsula; 16, Tuz Lake basin; 17, Malatya basin; 18, Kahta-Adiyaman; 19,

Yayladağ-Antakya.

The faunal succession of the upper part of the

section shows an alternation of low diversity and

high diversity assemblages of rudists. Several

dozen specimens of Pseudopolyconites have been

found and more than 90% of them have been

recovered in a small area (Liljekar) where a rich

monospecific assemblage is present. The

individuals are mainly large in size, always

isolated, separated from one another, and mostly in

grwoth position and often fully articulated. The

finding of this rudist outside this area is remarkably

rare.

It is not easy to explain the reason for the

abundance of specimens of Pseudopolyconites in

this small area. Due to the poor quality of the

exposures, both lithology and sedimentology are

not much help and the interpretation of the

depositional setting is questionable. It is speculated

that Pseudopolyconites individuals grew in

unconsolidated sediments on silty substrates in a

shallow shelf environment. The sediment included

between the spines of Pseudopolyconites consists

of ochreous clayey-silt suggesting that these rudists

thrived in muddy, low energy settings. The rudists

herein were presumably adapted to a life in turbid,

depositional environments. The life of individual

rudists was probably terminated suddenly by rapid

burial by sediments during a single sedimentation

episode (Tarlao et al., 2010).

3. RELEVANT CHARACTERISTICS OF

PSEUDOPOLYCONITES AND THE QUESTION OF THE

PSEUDOPOLYCONITES SPECIES INSTITUTED AT

BAČEVICA

All the examples of Pseudopolyconites recovered at

Bačevica show a dense coat of tubules or spines

(Figure 2a) emerging from the shell wall and then

curving downwards (Figure 2b) (Milovanović,

1937). This character is rarely found in other

localities where Pseudopolyconites has been

recovered. The tubules at the base are thin and short

but they became progressively larger and longer

during the growth of individuals. The thickness of

the wall of the tubules is large in comparison with

the diameter of the internal hole which is narrow.

The origin of the spines in Pseudopolyconites

was explained by Pons and Vicens (2008) by

examining the outer shell layer constructional

pattern which is considered the main diagnostic

character of the Radiolitidae since its pattern is used

both in the phylogeny and taxonomy of the family.


109

Figure 2. a) Transverse section of a right valve of a Pseudopolyconites serbicus specimen from Liljekar (Serbia);

b) right valve of a Pseudopolyconites serbicus individual showing the dense coat of spines emerging from the shell

wall; c) entire specimen of Pseudopolyconites serbicus from Liljekar; d) View from high of the left valve of the

specimen shown in c).

A transverse section and a radial section of an

example of Pseudopolyconites collected at

Bačevica are shown respectively in Figures 3-4.

The repetitive closed invaginations of the radial

downfolds in the growth lamellae, projecting

outwards as spines (Figure 4), is the main

characteristic of the outer shell layer structure of

Pseudopolyconites and this represents the extreme

case of the radial folding of the growth lamellae in

radiolitid rudists (Pons and Vicens, 2008).

However, one of us (Rajka Radoičić)

hypothesizes that the tubules are not integral parts

of the shell and may represent epizoa. Besides,

Pseudopolyconites tubules were explained by

Misik (1966, pl. LXXV, fig. 1) as serpulid worm

tubules (Filigrana sp.) from the serpula-bioherm

limestones of the Brezova Mountains (Western

Carpathians, Slovakia).

The Serbian rudistologists erected numerous

species of Pseudopolyconites in the environs of

Bačevica. The Branislav Milovanović old collection

of rudists included the five holotypes of the first

Pseudopolyconites species erected by Milovanović

(1934a, 1935a) and it was housed at the Institute of

Paleontology (Faculty of Philosophy) of the the

Belgrad University: it was lost during World War II

when the building was set on fire. Unfortunately,

the holotypes of the species established successively

by Milovanović and Sladić (1957) and by Sladić-

Trifunović (1986) have also vanished.

The most significant criterion selected by the

Serbian workers for establishing different species of

Pseudopolyconites was founded on the

characteristics of the ligamental ridge in the

transverse section (e.g., shape, length and

thickness). Minor characteristics concerning the


110

Figure 3. Spines seen in transverse section of the right

valve of a Pseudopolyconites serbicus specimen

(Liljekar).

Figure 4. Vertical section of the right valve of a

Pseudopolyconites serbicus specimen (Liljekar)

showing the outwards expansion of the spines (white

line). Shell wall on the right.

radial structures and the ornamentation have been

observed and discussed mainly by Sladić-Trifunović

(1983). Moreover, according to this author, the

shape of the ligamental ridge of Pseudopolyconites

individuals was remarkably transformed through

ontogenetic development of the right valve, and this

was the main criterion used for discrimining

species.

Tarlao et al. (2010) have considered all species

of Pseudopolyconites erected at Bačevica as simple

eco-morphotypes which may be attributed to just

one species, Pseudopolyconites serbicus, the earliest

one established by Milovanović (1934a).

It is stressed here that the level rich in examples

of Pseudopolyconites is included within a thin

fossiliferous lithosome and occupies a precise

stratigraphic position (i.e., Liljekar) along the

Bačevica section. We hypothesize also that this

situation was not different at the time the original

research was carried out by Milovanović and

Sladić-Trifunović as they often mention the place

name of Liljekar where we have found the majority

of the specimens of Pseudopolyconites.

4. PSEUDOPOLYCONITES BEARING STRATA WITHIN

THE CARBONATE SUCCESSIONS OF THE CENTRAL

TETHYS

Examples of Pseudopolyconites have been found

within late Campanian carbonate successions

cropping out in the northern Adriatic region (NE

Italy, Slovenia, Croatia, Bosnia), in southwestern

Italy and western Greece (Figure 1). Other species

of Pseudopolyconites were erected on the basis of

the usual criterion, that is, the change of the shape

of the ligamental ridge. Pseudopolyconites ovalis

apuliensis was described from Poggiardo (Apulia,

southern Italy) by Sladić-Trifunović and

Campobasso (1980) and Pseudopolyconites

campobassoi was described from Pokonj Dol (Hvar,

Croatia) by Sladić-Trifunović (1980).

The Upper Cretaceous limestones exposed on

the islands of Hvar and Brač typify the Adriatic

carbonate platform system. Pseudopolyconites

individuals, together with other radiolitids, are

embedded in massive bioclastic packstone to

rudstone of the Brač Marbles Unit of Pučišća

Formation. The foraminiferal association of

Orbitoides tissoti and Pseudosiderolites vidali is

referred to the middle or early late Campanian by

Gušić and Jelaska (1990) and was recalibrated by

strontium isotope stratigraphy to Middle Campanian

by Steuber et al. (2005).

Large fragments of massive valves of

Pseudopolyconites were collected within bioclastic

deposits at Mt. Nanos, Western Slovenia (Pleničar,


111

2005), Mt. Jouf, NE Italy (Sladić-Trifunović and

Nereo, 1990) and, more recently from an olistolith

at Anhovo, northwestern Slovenia (Pleničar and

Jurkovšek, 2009). Pseudopolyconites manjae

Milovanovic & Sladic was erected on material

from Mt. Jouf. Numerical ages derived from the

strontium isotope stratigraphy indicated, in general,

a late Campanian age for the Pseudopolyconites

bearing strata at Mt. Jouf (Swimburne and Noacco,

1993). Pseudopolyconites slovenicus Pleničar &

Jurkovšek recovered from Anhovo is the latest

species added to the conspicuous catalog of the

genus in question.

The rudist shell beds of all the previously

mentioned localities from Croatia to Apulia can be

considered hydraulic shell concentrations which

were deposited under the influence of hydraulic

processes with the input of surrounding bioclastic

sediments. Thus, due to the hard lithology, it is

difficult to collect good examples of

Pseudopolyconites.

Recently, late Cretaceous platform carbonate

successions containing specimens of

Pseudopolyconites from the Salento peninsula

(southern Italy) were studied by strontium isotope

stratigraphy (Schlüter et al., 2008a). The rudist

associations of the S. Cesarea Limestone exposed

in the quarry near Vitagliano (late Campanian) and

those of the overlying Ciolo Limestone exposed in

the Cava Soframa and Ciolo cove (late

Maastrichtian), from a time interval of more than

10 Ma years, are surprisingly similar. Successively

(Steuber et al., 2007), numerical ages of well

preserved rudist shells from breccias and

megabreccias of the slope of the same Apulian

Platform exposed on the Ionian islands (W. Greece)

also indicate a late Maastrichtian age. Some

characteristic genera, such as, Joufia,

Pseudosabinia and other caniculate recumbents, are

abundant throughout this time interval both in the

Salento peninsula and in the Ionian islands. Many

of these rudists (Pseudopolyconites sp., Hippurites

cornucopiae, Plagioptychus sp., Pironaea polystila,

Joufia reticulata, Pseudosabinia sp., Mitrocaprina

sp., etc.) are also present in the clastic successions

cropping out in the environs of Bačevica, Gavlo

(Bulgaria) and in some localities of Turkey.

5. PRESENCE OF PSEUDOPOLYCONITES-BEARING

STRATA WITHIN CLASTIC SUCCESSIONS OF

SOUTHEASTERN EUROPE, TURKEY AND WITHIN

CARBONATE SUCCESSIONS OF THE MIDDLE EAST

Rudist shells are abundant in the Campanian and

Maastrichtian rudist-bearing deposits of the Peri-

Adriatic intra-oceanic carbonate platforms, but they

are usually reworked and broken and often beyond

taxonomic recognition.

Examples of Pseudopolyconites have been

found within mixed carbonate-siliciclastic or

volcanoclastic Campanian successions (Figure 1) in

Eastern Serbia (Rtanj and Vrla Strana, Vina),

Bulgaria (Gavlo), Rumania (Magura Hill and

Remeti valley) and Turkey (Kocaeli peninsula-

Istanbul zone, Tuz Lake-Central Anatolia, Malatya-

Eastern Anatolia, Adiyaman and Antakya-

Southeastern Anatolia). Rudists here are often well

preserved, isolated and in growth position.

The Bulgarian outcrops at Garlo (Breznik area)

are about 120 km from those of Bačevica-Vrbovac

and are closely related: both sequences are

underlain by lavas and tuffs and consist mainly of

sandstones and siltstones with subordinate

conglomerates, marlstones and limestones.

Excluding the plethora of rudist species, concerning

in particular the genera Pironaea and Biradiolites,

studied by Milovanović (1934b and 1935b) and

Pamouktchiev (1964, 1975, 1979) respectively, both

the Bačevica and Garlo successions have very

similar rudist faunas. Given these strong similarities

it is supposed that the age of the deposits in eastern

Serbia and western Bulgaria is similar. The age of

the Pironaea-Joufia-Pseudopolyconites assemblage

at Garlo, as determined from strontium isotopes is

late Campanian (Swinburne et al., 1992).

The finding of Pseudopolyconites specimens is

rare at Garlo. A new species Pseudopolyconites

garlensis was established by Pamouktchiev (1979).

But the poor description and illustrations of this

form mean that we cannot form an opinion on the

validity of this species.

Late Santonian-early Maastrichtian(?) rudist-

bearing strata widely crop out on the Apuseni

Mountains (Rumania). The Apuseni Mountains

comprise a sedimentary succession comparable to

the well-studied Gosau Group in the Eastern Alps

and in other regions of the Alpine-Carpathians chain

(Western Carpathians and Transdanubian range).

The estimates of the ages of these deposits rely on

nannofossils, benthic foraminifers and rudist

bivalve biostratigraphy but not on strontium isotope

stratigraphy.

Pseudopolyconites is a rare rudist bivalve in

Rumania. It was found only in two localities of

Apuseni Mountains: Magura Hill

(Pseudopolyconites hirsutus, Patrulius, 1974;

Pseudopolyconites parvus, Săsăran et al., 2013) and

Remeti Valley (Pseudopolyconites milovanovici,

Lupu, 1969, 1974).

A good example of Pseudopolyconites was

found in Hron Valley, Slovakia (Mišik, 1966; Lupu,

1976).


112

Specimens of Pseudopolyconites have been

recovered in late Campanian-Maastrichtian

transgressive mixed siliciclastic-carbonate

successions of Turkey, from the Istanbul zone

(Kocaeli peninsula, northwestern Turkey), Tuz

Lake Basin (central Anatolia), Malatya Basin

(eastern Anatolia) to Adıyaman and Antakya areas

(southeastern Turkey) but they are not abundantly

represented (Özer, 1982, 1983, 1992 a, b, 2002;

Özer et al., 2008, 2009; Steuber et al., 2009).

However, most examples of Pseudopolyconites are

well preserved, isolated, generally large, in growth

position and free of sediment matrix which makes

it possible to observe all the internal and external

characteristics of both valves.

The Hereke area (Kocaeli peninsula) is the type

locality of Pseudosabinia, where some

Pseudopolyconites specimens are also reported as

occurring together with Gorjanovicias by Özer

(1982, 1992a) and Fenerci (1999). The previous

studies suggest a Campanian, Campanian-

Maastrichtian or Maastrichtian age for the rudist

fauna, however the Sr-isotope data of the rudist

shells indicate a late Campanian age for the rudist-

bearing limestones in the Hereke area (personal

communication with T. Steuber).

The transgressive sequence of Tuz Lake basin

contains well-preserved specimens of

Pseudopolyconites (Özer, 1983) together with

abundant hippuritids (Özer, 2002). The Sr-isotope

analysis of the equivalent levels of the rudist-

bearing limestones in the north of the basin, around

Ankara, indicate a late Campanian age (personal

communication with T. Steuber).

Rudists are widespread and very abundant in

the Campanian-Maastrichtian transgressive-

regressive systems tracts of the Malatya Basin

(Özer et al., 2008). In the southern part of this

basin, around Yeşilyurt village, Sr-isotope data

obtained from rudists of the Pseudopolyconites-

bearing limestones just below the Orbitoides

apiculatus level indicate a late Campanian age

(Özer et al., 2008; Schlüter et al., 2008a; Schlüter,

2008b).

The Kahta-Adıyaman and Yayladağı-Antakya

areas are the type localities of the several rudist

taxa that are endemic to the Arabian platform-plate

(Özer, 1986, 1992b; Steuber et al., 2009). The

rudist-bearing limestone lenses within the clastic

sequence contain some specimens of

Pseudopolyconites (Özer, 1986, 1991; Özer et al.,

2008). Strontium isotope analysis of rudist shells

indicates a late Campanian age for the

Pseudopolyconites-bearing limestone lenses in the

transgressive sequence in the Kahta and Yayladağı

areas (Özer et al., 2008; Schülter et al., 2008b;

Steuber et al., 2009).

As far as the Middle East is concerned, the best

outcrops of late Campanian-Maastrichtian rudist

bearing sequences are in Iran (Khazaei et al., 2010

with bibliography) but Pseudopolyconites

specimens have not been found so far in Iran

(Khazaei and Özer, 2011). Rudist collections from

NW and NE Syria with similar rudist assemblages

to those of southeastern Anatolia have been recently

examined by one of us (S. Ö.) indicating a late

Campanian-early Maastrichtian age. But,

unfortunately, Pseudopolyconites has not been

found. The rudists are very limited in the Upper

Cretaceous sequences of Jordan and they are only

observed in the Cenomanian and Turonian beds

(Bandel and Mustafa, 1994; Özer and Ahmad,

2011) indicating that it seems impossible to find

Pseudopolyconites specimens. The late Campanian-

Maastrichtian rudist fauna of NE Iraq showing close

resemblances with those of southeastern Anatolia

and Iran have been recently presented by Özer et al.

(2012), but Pseudopolyconites specimens have not

been observed.

Our knowledge of the rudist fauna of the Middle

East has been improved recently by Steuber and

Schlűter (2012) who presented a strontium-isotope

stratigraphy of Upper Cretaceous rudist bivalves of

the Arabian Plate. Due to the presence of

Pseudopolyconites in southeastern Anatolia (Özer,

1986, 1991, 1992b; Steuber et al., 2009) and in the

United Arab Emirates (Morris and Skelton, 1995),

the existence of this taxon in other localities of the

Middle East is highly probable.

6. ANCESTORS OF THE CAMPANIAN-

MAASTRICHTIAN PSEUDOPOLYCONITES SPECIES

The oldest representative of Pseudopolyconites is

considered to be Pseudopolyconites hirsutus

(=Duranddelgaia hirsuta) Patrulius, 1974,

described from the Apuseni Mountains, Romania.

Pejovic and Sladić-Trifunovic (1977) were the first

authors who observed that the specimens described

as Duranddelgaia hirsuta (Patrulius, 1974) have all

the specific characters of the genus

Pseudopolyconites and so there was no need to

establish the new genus Duranddelgaia. Specimens

of P. hirsutus (Patrulius) from Măgura Hill

(Apuseni Mountains) are represented by the

holotype, two paratypes and six specimens that are

housed at the Geological Museum in Bucarest

(Figure 5). Of these six specimens, four are poorly

preserved right valves and the two others are large

fragments of the right valve embedded in rock.

Previously, P. hirsutus (Patrulius) was considered to

be an Early Santonian representative of


113

Figure 5. a) Holotype P11060 of Pseudopolyconites

hirsutus: dorsal side of right valve with tubular

excrescences; b) specimen 11063A of

Pseudopolyconites hirsutus: right valve with ventral

radial band. Original specimens collected by

Patrulius (1974) are in the custody of the Geological

Museum in Bucharest (Rumania).

Pseudopolyconites based on Patrulius (1974).

Recently, new detailed investigations on the rudist

fauna from Măgura Hill have been made and

independent biostratigraphic data (e.g.,

nannoplankton) as well as the rudist assemblages

suggest a Late Santonian-Early Campanian age

(Săsăran et al., 2013). Additionally, new rudist taxa

(Pseudopolyconites parvus and Pseudosabinia

klinghardti) were described for the first time from

these deposits (Săsăran et al., 2013).

The specimens of P. hirsutus and P. parvus

described from Măgura Hill by Săsăran et al.

(2013) have smaller dimensions as compared to

specimens describes from Serbia (Bačevica, Lesak)

(Pejovic and Sladić-Trifunovic, 1977; Sladić-

Trifunovic, 1983) and have no long spines. Also,

detailed taxonomical investigations revealed an

intraspecific variability of external morphological

features of both valves of P. hirsutus as follows:

the shape of left valves varies from flat and very

thin to slightly convex and thicker while the shape

of right valves varies from conical to cylindrical-

conical, slightly curved in some specimens. Also, in

the right valve the ventral radial band is always flat,

without tubular excrescences while the posterior

radial band usually is not discernable on the surface

of the, but may appear as a narrow furrow with one

fine, longitudinal costa in some specimens (Săsăran

et al., 2013). The same variability of external

morphological features of both valves in small

species of Pseudopolyconites has been also

observed by Sladić-Trifunovic (2004) within the

rudist fauna from Lesak (Serbia).

Pseudopolyconites hirsutus and P. parvus have

been found in mixed siliciclastic-carbonate deposits

at Măgura Hill that transgressively overlie

Mesozoic tectonic units (Codru Nappe complex).

The stratigraphic succession is 40 m-thick and starts

with basal conglomerates which are succeeded by

sandstone/marlstone intercalations with plant

remains and mollusc fragments. These are followed

by gastropod-rich limestones dominated by

actaeonellids and nerineids, with rare rudists. These

limestones are overlain by marls and marly

limestones rich in solitary and meandroid colonial

corals. The succession continues with interlayers of

marls/sandstones/conglomerates containing three

calcareous levels (Săsăran et al., 2013).

Specimens of P. hirsutus were also recognized

by Pejović and Sladić-Trifunović (1977) in a large

olistolith characterized by abundant rudist fauna at

Svračija Stena, Lešak (southwestern Serbia).

According to these authors, the forms of P. hirsutus

recovered at Lešak are identical with the presumed

coeval rudist bivalves from Măgura Hill. Based on

species of Sulcoperculina aff. cubensis (Palmer)

Radoičić ascribes the Lešak olistholit to Campanian

or Upper Santonian-Campanian.

Successively, Sladić-Trifunović (1983)

established the new species P. leposavicensis from

Svračija Stena and in her last paper (2004) she

instituted Pseudopolyconites pantici in place of P.

hirsutus and P. leposavicensis. We find it

impossible to understand the reason for the

emendation by changing the name of P. hirsutus

made by Sladić-Trifunović (1983; 2004) because

we consider that the two mentioned species of P.

leposavicensis and P. pantici are synonymous with

P. hirsutus falling into intraspecific variability of

this species. P. parvus was previously known only

from Late Campanian sequences from Eastern

Serbia (Bačevica).

7. DISCUSSION

Examples of Pseudopolyconites were found mostly


114

within the late Campanian successions. They have

been found in many localities but they are almost

never abundant except close to Bačevica. In

particular, owing to reworking in high energy

environments typical of the Central Tethys

carbonate platforms during the Campanian,

complete specimens of Pseudopolyconites are very

rare and, often, only fragmented shells have been

collected. Moreover, the outer shell layer characters

of Pseudopolyconites established far from Bačevica

are partly preserved or, in some cases, quite lacking

because they were removed during the mechanical

breakdown in the high-energy environments. Only

photos of poorly preserved Pseudopolyconites

and/or large fragments of the right valves of the

same in addition to cross sections are available in

most cases. We stress that, in spite of long and

laborious research carried out in Belgrad, the

original palaeontological material with all the

holotypes and paratypes of the species of

Pseudopolyconites established by the Serbian

rudistologists both in Serbia and abroad has not

been found, making comparative analysis between

the holotypes and topotypes impossible. To sum

up, it is difficult to hold for certain the validity of

all the species of Pseudopolyconites established by

the various authors (i.e., P. campobassoi, P. ovalis

apuliensis, P. manjae, but also P. garlensis, P.

milovanovici and P. slovenicus). Further, the

numerous ‘species’ of Pseudopolyconites from

Bačevica have been considered as simple eco-

morphotypes (Tarlao et al., 2010). The proliferation

of species made by rudistologists at Bačevica is not

only peculiar to the genus Pseudopolyconites. For

instance, three different ‘species’ of Pironaea

established by Milovanović (1934b) have been

recovered by us within the same fossiliferous level

(see also Swimburne et al., 1992 and Munujos

Vinyoles, 1989, for discussion about the

determination of Pironaea species).

Pseudopolyconites was not present only during

the late Campanian since specimens have been

recovered in other Late Cretaceous successions of

Central Tethys-Vardar zone. The earliest forms of

Pseudopolyconites were recovered from Dealul

Magura (Patrulius, 1974) and from the Lešak

olistolith (Pejović and Sladić-Trifunović, 1977) and

were considered to be the same species. In the

absence of age diagnostic fossils and of strontium-

isotope analysis, a precise chronological attribution

of the deposits cropping out in these two localities

is not yet established. However, the rudist taxa of

Magura Hill and other biostratigraphic data (e.g.,

nanoplankton) suggest a late Santonian-early

Campanian age (Săsăran et al., 2013). All the

original examples from Lešak seem to have

vanished and, moreover, the Serbian locality is at

present unreachable. It is just on the boundary

between Kosovo-Metohia region and Serbia. Only

photos of selected transverse sections of

Pseudopolyconites from Lešak but fine entire

examples from Magura Hill are available. A few

slight differences concerning mainly the different

shape of the tip of the ligamental ridge were

observed by Sladić-Trifunović (2004). Furthermore,

the individuals from Rumania seem to be larger in

size in comparison with the Serbian ones. To sum

up the synonymy of these two old forms of

Pseudopolyconites is probable. Transitional forms

of Pseudopolyconites from Santonian to late

Campanian are not known. An “evolutionary

phylogenetic jump” (sensu Sladić-Trifunović, 2004)

characterized mostly by size increase and by the

development of a rich coat of tubules.

Pseudopolyconites individuals are relatively

abundant in the late Campanian deposits: the so-

called specific Pironaea-Pseudopolyconites

association, widely distributed in the Central

Tethys, represents an important palaeo-

biogeographic and biostratigraphic event in the area

in question (Milovanović and Grubić, 1971; Sladić-

Trifunović, 1983). Following the datum that the sea-

water Sr/Ca had reached a particularly high value in

the Campanian (Stanley and Hardie, 1998; Steuber

and Veizer, 2002) and coeval rudist shells were

calcite-dominated (Steuber, 2002), it may be that

Pseudopolyconites shells benefitted by the relative

abundance of calcite by developing very much their

growth lamellae structure. This may be one of the

possible reasons for the extraordinary growth of

tubules in late Campanian Pseudopolyconites.

Another hypothesis is that Pseudopolyconites which

lived in siliciclastic (or vulcanoclastic) settings

developed peculiar functional adaptations of the

shells different from the specimens which thrived in

carbonate settings (see also Steuber, 1997).

Pseudopolyconites representatives undoubtedly

attributed to the early Maastrichtian are not yet

known. Pseudopolyconites examples have been

recovered within rudist-bearing deposits dated back

to the latest Maastrichtian of the Apulian carbonate

platform (Steuber et al., 2007; Schlüter et al.,

2008a). Unfortunately the Pseudopolyconites

examples recognized within these deposits are

poorly preserved and the collection of complete

shells or larger fragments for detailed descriptions is

almost impossible. It would be very important to

take fine examples from the hard rock to evaluate

possible differences between late Campanian and

latest Maastrichtian Pseudopolyconites. The

palaeontological data presented by Steuber et al.

(2007) and Schlüter et al. (2008a) provide evidence


115

that the rudist associations existing in the latest

Maastrichtian, close to the K-P boundary of the

Apulian carbonate platform are remarkably similar

to the rudist assemblages recognized within the late

Campanian limestones of the same platform. The

rudist associations of Bačevica and of the nearby

locality of Vrbovac, characterized by a conspicuous

group of caniculate rudists, are also surprisingly

similar to those of the Apulian Plate exposed in

Salento (S. Italy) and in the Ionian islands

(Greece).

Because of the superb preservation of

Pseudopolyconites our conjectures concerning the

palaeoecology are entirely based on the Bačevica

Pseudopolyconites-rich site (i.e., Liljekar). It is

observed that both the short and long spines reach

the base of the lower valve in Pseudopolyconites. It

is speculated that the spines were useful for

attaching the shells to the substrate. The spines

emerging from the outer shells of P. hirsutus seem

to curve upwards. In that case, they probably had a

different function in comparison with those of P.

serbicus, maybe acting as a defense against

possible predators.

Pseudopolyconites from Bačevica thrived in

moderate to low energy settings, maybe in, or close

to, shallow channels and creeks of muddy tidal

flats. It is hypothesized that the group of caniculate

rudists found within the rudist-bearing lithosomes

of Bačevica and Vrbovac colonized intertidal zones

of protected marine environments (Özer and

Ahmad, 2011). Shallow lagoons and/or tidal flats

are sufficiently represented both in the carbonate

and siliciclastic Central Tethys realm during the

late Campanian. But, due to the general drowning

and, sometimes, demise of the carbonate platform

system in the Central Tethys during the

Maastichtian, tidal flat environments are extremely

rare in the early Maastrichtian seas of the Central

Tethys and scarcely documented during the late

Maastrichtian (Dercourt et al., 1993). This may

explain the relative rarity of Pseudopolyconites

during the Maastrichtian.

Lastly, as far as the taxonomy of

Pseudopolyconites is concerned, Sladić-Trifunović

(1983) proposed the segregation from radiolitids as

a new family (i.e., Pseudopolyconitidae) for the

genera Pseudopolyconites, Fundinia (Sladić-

Trifunović and Pejovic, 1977) and Kurtinia

(Karacabay-Öztemur, 1980) based on the outer

shell layer structure are the growth lamellae, which

project outwards as spines. Pons and Vicens (2008)

showed that this character alone may not be used to

define monophyletic groups. Moreover, Kurtinia

cannot be ascribed to Pseudopolyconites. Fundinia

also cannot be regarded as a member of the

“pseudo-family”. In fact Fundinia and Kurtinia

have no spines emerging from the shell wall which

are the most remarkable elements in the structure of

Pseudopolyconites.

The new updated phylogenetic classification of

rudist bivalves proposed by Skelton (2013)

definitively obliterates the family of

Pseudopolyconitidae.

8. CONCLUSIONS

Pseudopolyconites can be considered a rare

component of Upper Cretaceous faunal

assemblages. Many late Campanian species of

Pseudopolyconites were instituted in the environs of

Bačevica and elsewhere. Many species were based

on poorly preserved specimens and, often using

cross sections. This makes it difficult or even

impossible to evaluate all the characters necessary

for an unequivocal separation amidst the previously

established species.

Thus, at the present state of knowledge, many

specimens of Pseudopolyconites previously

assigned to different species might fall within the

biological variability of a single species. A few

characters allow us to separate some late Santonian-

early Campanian representatives of the genus from

the late Campanian ones, but it is not possible to

form an opinion concerning poorly preserved

Pseudopolyconites from the latest Maastrichtian. It

may be that Pseudopolyconites preserved unaltered

the peculiar characters from the late Campanian up

to the latest Maastrichtian, analogously to a

conspicuous group of caniculate rudist which seem

to have passed the fore-mentioned long lapse of

time without significant changes of the skeletal

characters. Thus, the topic of functional adaptations,

growth rates and different shell growth strategies of

Pseudopolyconites from the Santonian to the latest

Maastrichtian should be adequately examined in the

future, considering also the different environments

where these radiolitids thrived, for instance,

carbonate versus siliciclastic/volcanoclastic settings.

Specimens of Pseudopolyconites found at

Bačevica are considered herein as rudist bivalves

which developed particular shell growth strategies

in response to the peculiar environments where they

thrived, probably in muddy tidal flats.

This paper may underestimate the true richness

of Pseudopolyconites, but without more well-

preserved material this cannot be considered.

Undoubtedly phylogenetic relationships within the

genus in question are obscure and a future job

concerning accurate descriptions and character

analysis both of historical and new examples will be

needed.


116

Acknowledgements. This paper has benefited from the

careful reviews by Simon Mitchell and one anonymous.

We are most grateful to Josè Maria Pons who read our

paper at the 9th International Congress on Rudist bivalves

in Kingston, Jamaica in 2011.

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Accepted 21st June 2013

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